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Ün0£5Teicreo ANTIGENIC VARIATION in VIRULENCE Open Research Online The Open University’s repository of research publications and other research outputs Antigenic Variation in Virulence Determinants of Streptococcus zooepidemicus and Actinobacillus equuli Involved in Lower Airway Disease of the Horse and Strategies Towards Protective Immunisation Thesis How to cite: Ward, Chantelle Louise (1998). Antigenic Variation in Virulence Determinants of Streptococcus zooepidemicus and Actinobacillus equuli Involved in Lower Airway Disease of the Horse and Strategies Towards Protective Immunisation. PhD thesis The Open University. For guidance on citations see FAQs. c 1997 Chantelle Louise Ward https://creativecommons.org/licenses/by-nc-nd/4.0/ Version: Version of Record Link(s) to article on publisher’s website: http://dx.doi.org/doi:10.21954/ou.ro.0000f95a Copyright and Moral Rights for the articles on this site are retained by the individual authors and/or other copyright owners. For more information on Open Research Online’s data policy on reuse of materials please consult the policies page. oro.open.ac.uk üN0£5Teicreo ANTIGENIC VARIATION IN VIRULENCE DETERMINANTS OF STREPTOCOCCUS ZOOEPIDEMICUS AND ACTINOBACILLUS EOUULI INVOLVED IN LOWER AIRWAY DISEASE OF THE HORSE AND STRATEGIES TOWARDS PROTECTIVE IMMUNISATION BY CHANTELLE LOUISE WARD, BSc A thesis submitted in partial fulfilment of the requirements of the Open University for the degree of Doctor of Philosophy, pertaining to the discipline of microbiology December 1997 Animal Health Trust P.O. Box 5 Newmarket Suffolk CB8 7DW Work supported by Hoechst Roussel Vet. Ltd. Offie OP mwfw : '27 s’ytw ProQuest Number:C804782 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest ProQuest C804782 Published by ProQuest LLO(2019). Copyright of the Dissertationis held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code Microform Edition © ProQuest LLO. ProQuest LLO. 789 East Eisenhower Parkway P.Q. Box 1346 Ann Arbor, Ml 4 8 1 0 6 - 1346 ABSTRACT Lower airway disease (LAD) of the training Thoroughbred horse is characterised by poor performance and excess mucus and pus in the trachea. This condition, although often referred to as ‘the virus’, is significantly associated with Streptococcus zooepidemicus and Actinobacillus/Pasteurella species without evidence of viral infection. The degree of antigenic variability in these bacteria was assessed, to judge the number of strains that might be required in future vaccines. Particular attention was paid to • • • the putative virulence determinants of the M-like protein and capsule of S. zooepidemicus and the iron regulated proteins (IRPs), produced only in low available iron concentrations, of Actinobacillus equuli. ' * ' , ■ -V-!- A range of Actinobacillus/Pasteurella species were isolated from the lower airway of horses, but more than half were accounted for by A.equuli. A. equuli bound and utilised equine holotransferrin for growth. The constitutive proteins of A. equuli were antigenically cross-reactive, but IRPs were more antigenically varied. Vaccines based on interference of IRPs would be likely to need more than one strain of A. equuli. Hot acid extracts of S. zooepidemicus were antigenically cross-reactive, but there was less opsonogenic cross-reactivity in horse blood containing natural antibodies than rabbit antisera to extracts. A vaccine based on M-like protein would be likely to need more thah one type of S. zooepidemicus, but no relationship could be found between ribotype / antigenic profile and opsonogenic behaviour, suggesting that M-like protein is not necessarily the principal anti-phagocytic mechanism for these isolates, at least in vitro. • s. zooepidemicus produces a hyaluronate capsule thought to be a virulence determinant, which is tightly bound by a protein (HAP). Immunisation with recombinant HAP significantly increased survival time and clearance of bacteria in mice challenged intraperitoneally with S. zooepidemicus. Recombinant HAP may have acted as a surrogate antigen for the capsule. Introduction of bacteria into ponies intratracheally produced a dose response for S. zooepidemicus and A. equuli. However, so much intercurrent infection developed (due to S. zooepidemicus and Bordetella bronchiseptica) it was concluded that conventional ponies would be unsuitable for sttaightforward studies of vaccine efficacy in response to experimental challenge. Alternative ways forward are discussed. Ward, C. 1997 SUMMARY Lower airway disease (LAD) of the training Thoroughbred horse is characterized by poor performance and excess mucus and pus in the trachea. This condition, although frequently referred to as “the virus”, is significantly associated with Streptococcus zooepidemicus and Actinobacillus/Pasteurella species without evidence of viral infection. The objective was to assess the degree of antigenic variability in these bacteria to judge the number of strains that might be required in future vaccines. Particular attention was paid to the putative virulence determinants of the M-like protein and capsule of S. zooepidemicus and the iron regulated proteins (IRPs), produced only in low available iron concentrations, of Actinobacillus equuli. More than half (52.1%) of the tracheal wash isolates originally characterized as Actinobacillus/Pasteurella species involved were identified as A. equuli, 18.3% were Actinobacillus suis -like, 12.7% were Pasteurella pneumotropica, 8.5% were Actinobacillus ligniersii , 7.0% were Pasteurella haemolytica and 5.6% were P. mairii, suggesting that a range of Actinobacillus/Pasteurella species can be isolated from the lower airway of horses, but more than half are accounted for by A.equuli. All isolates cultured from cases of LAD were selected on the basis of their temporal and geographical separation. In iron limited media, all isolates bound and utilised equine holotransferrin for growth, but not iron bound to rabbit transferrin, human transferrin, or a number of other equine iron carrying proteins. The constitutive proteins of A. equuli were antigenically cross-reactive, but IRPs were more antigenically varied. Vaccines based on interference of IRPs would be likely to need more than one strain of A. equuli. Hot acid extracts of S. zooepidemicus were antigenically cross-reactive, but there was less opsonogenic cross-reactivity in horse blood containing natural antibodies than rabbit antisera to extracts. A vaccine based on M-like protein would be likely to need more than one type of S. zooepidemicus, but no relationship could be found between ribotype / Ward, C. 1997 antigenic profile and opsonogenic behaviour, suggesting that M-like protein is not necessarily the principal anti-phagocytic mechanism for these isolates, at least in vitro. S. zooepidemicus produces a hyaluronate capsule thought to be a virulence determinant, which is tightly bound by a protein (HAP). Immunisation with recombinant HAP significantly increased survival time and clearance of bacteria in mice challenged intraperitoneally with S. zooepidemicus. Recombinant HAP may have acted as a surrogate antigen for the capsule. Ponies were selected for challenge at the place of supply on the basis of their limited serum antibody to IRPs or M-like proteins for the strains under study. After transport to Home Office designated premises and following introduction of 10*°, 10’ and 10® cfu intratracheally, a dose response for S. zooepidemicus and A. equuli was observed in severity of clinical observations and gross pathology at post mortem examination, 3 or 4 days after infection. However, so much intercurrent infection developed (due to S. zooepidemicus and Bordetella bronchiseptica ) it was concluded that conventional ponies would be unsuitable for straighforward studies of vaccine efficacy in response to experimental challenge. Alternative ways forward are discussed. ward, u. 199/ CONTENTS Summary. 1 Contents. ^ List of Tables. 6 List of Figures. 8 Acknowledgements. 10 1.0 Chapter One - Preface, Introduction and Objectives. 11 1.1 Contents. 12 1.2 Preface to the Introduction. 13 1.3 Introduction. 14 1.3.1 The equine respiratory tract - in health. 14 1.3.1.1 Anatomy and physiology. 14 1.3.1.2 Airway defence mechanisms. 15 1.3.1.3 Cytology and bacteriology. 18 1.3.2 The equine respiratory tract - in disease. 19 1.3.2.1 Diagnosis. 19 1.3.2.2 Aetiology. 21 1.3.2.3 Immune mechanisms during bacterial infection 24 1.3.2.4 Treatment and prevention of infection / transmission 26 1.3.3 Streptococcus zooepidemicus and related bacterial pathogens. 27 1.3.3.1 Classification. 27 1.3.3.2 S. zooepidemicus disease in mammals. 28 1.3.3.3 Pathogenic determinants 29 1.3.3.4 Vaccination strategies. 33 1.3.4 Actinobacillus equuli and related pathogens. 36 1.3.4.1 Classification. 36 13 A.2 A. equuli and related disease in mammals. 37 1.3.4.3 Pathogenic determinants. 38 1.3.4.4 Vaccination strategies. 41 wara, L,. ivv/ 1.3.5 Justification for study. 43 1.4 Aims and objectives. 45 2.0 Chapter Two - Actinobacillus and Pasteurella species isolated from the distal trachea of horses with bacterial lower airway disease. 46 2.1 Contents. 47 2.1.1Iist of Tables. 47 2.2 Objectives. 48 2.3 Summary. 49 2.4 Materials and Methods. 50 2.5 Results and Discussion. 53 3.0 Chapter Three ■ Utilisation and binding of equine transferrin by Actinobacillus equuli and characterisation of its iron regulated proteins. 59 3.1 Contents. 3.1.1 list of Tables. 60 3.1.2 list of Figures. 60 3.2 Objectives. 62 3.3 Summary. 63 ; 3.4 Materials and Methods. 64 3.5 Results and Discussion. 70 4.0 Chapter Four - Antigenic relatedness of isolates of Streptococcus zooepidemicus from the equine trachea and development of a murine challenge model. 84 4.1 Contents. ^5 4.1.1 list of Tables. 85 4.1.2 list of Figures. 85 4.2 Objectives.
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